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1.  Oil-producing flowers within the Iridoideae (Iridaceae): evolutionary trends in the flowers of the New World genera 
Annals of Botany  2012;110(3):713-729.
Background and Aims
Oil-producing flowers related to oil-bee pollination are a major innovation in Neotropical and Mexican Iridaceae. In this study, phylogenetic relationships were investigated among a wide array of New World genera of the tribes Sisyrinchieae, Trimezieae and Tigridieae (Iridaceae: Iridoideae) and the evolution of floral glandular structures, which are predominantly trichomal elaiophores, was examined in relation to the diversification of New World Iridaceae.
Methods
Phylogenetic analyses based on seven molecular markers obtained from 97 species were conducted to produce the first extensive phylogeny of the New World tribes of subfamily Iridoideae. The resulting phylogenetic hypothesis was used to trace the evolutionary history of glandular structures present in the flowers of numerous species in each tribe. Hypotheses of differential diversification rates among lineages were also investigated using both topological and Binary-State Speciation and Extinction methods.
Key Results and Conclusions
Floral glandular structures and especially trichomal elaiophores evolved multiple times independently in the American tribes of Iridoideae. The distribution pattern of species displaying glandular trichomes across the phylogeny reveals lability in the pollination system and suggests that these structures may have played a significant role in the diversification of the Iridoideae on the American continent.
doi:10.1093/aob/mcs134
PMCID: PMC3400458  PMID: 22782239
Elaiophores; glandular trichomes; Iridoideae; nectaries; pollination systems; Sisyrinchieae; Tigridieae; Trimezieae
2.  Pollination Biology of Jacaranda oxyphylla with an Emphasis on Staminode Function 
Annals of Botany  2008;102(5):699-711.
Background and Aims
Bignoniaceae is a Neotropical family with >100 genera, only two of which, Jacaranda and Digomphia, have a developed staminode. Jacaranda oxyphylla, whose flowers possess a conspicuous glandular staminode, is a zoophilous cerrado species. Here, the composition of the secretion of the glandular trichome and the influence of the staminode on the pollination biology and reproductive success of J. oxyphylla were studied.
Methods
The floral morphology, pollen viability, stigma receptivity, nectar volume and nectar concentration were studied. Compatibility system experiments were performed and floral visitors were observed and identified. Experiments comparing the effect of staminode presence and absence on pollen removal and pollen deposition efficiency were conducted in open-pollinated flowers. Histochemistry, thin-layer chromatography (TLC) and gas chromatography coupled to flame ionization detection (GC–FID) analyses were performed to determine the main chemical components of the staminode's glandular trichome secretion.
Key Results
Flower anthesis lasted 2 d and, despite the low frequency of flower visitation, pollination seemed to be effected mainly by medium-sized Eulaema nigrita and Bombus morio bees, by the small bee Exomalopsis fulvofasciata and occasionally by hummingbirds. Small bees belonging to the genera Ceratina, Augochlora and Trigona were frequent visitors, collecting pollen. Jacaranda oxyphylla is predominantly allogamous. Staminode removal resulted in fewer pollen grains deposited on stigmas but did not affect total pollen removal. The secretion of capitate glandular trichome occurs continually; the main chemical compounds detected histochemically were phenolic and terpenoid (essential oils and resins). Monoterpene cineole, pentacyclic triterpenes and steroids were identified by TLC and GC–FID.
Conclusions
The staminode of J. oxyphyllla is multifunctional and its importance for female reproductive success was attributed mainly to the secretion produced by capitate glandular trichomes. This secretion is involved in complex chemical interactions with pollinating bees, including the solitary bees Euglossini. These bees are common pollinators of various species of Jacaranda.
doi:10.1093/aob/mcn152
PMCID: PMC2712375  PMID: 18765441
Bignoniaceae; Jacaranda oxyphylla; pollination; bee; staminode; glandular trichomes; reproductive success; terpenes; steroids; phenolics
3.  Radiation of Pollination Systems in the Iridaceae of sub-Saharan Africa 
Annals of Botany  2006;97(3):317-344.
• Background Seventeen distinct pollination systems are known for genera of sub-Saharan African Iridaceae and recurrent shifts in pollination system have evolved in those with ten or more species. Pollination by long-tongued anthophorine bees foraging for nectar and coincidentally acquiring pollen on some part of their bodies is the inferred ancestral pollination strategy for most genera of the large subfamilies Iridoideae and Crocoideae and may be ancestral for the latter. Derived strategies include pollination by long-proboscid flies, large butterflies, night-flying hovering and settling moths, hopliine beetles and sunbirds. Bee pollination is diverse, with active pollen collection by female bees occurring in several genera, vibratile systems in a few and non-volatile oil as a reward in one species. Long-proboscid fly pollination, which is apparently restricted to southern Africa, includes four separate syndromes using different sets of flies and plant species in different parts of the subcontinent. Small numbers of species use bibionid flies, short-proboscid flies or wasps for their pollination; only about 2 % of species use multiple pollinators and can be described as generalists.
• Scope Using pollination observations for 375 species and based on repeated patterns of floral attractants and rewards, we infer pollination mechanisms for an additional 610 species. Matching pollination system to phylogeny or what is known about species relationships based on shared derived features, we infer repeated shifts in pollination system in some genera, as frequently as one shift for every five or six species of southern African Babiana or Gladiolus. Specialized systems using pollinators of one pollination group, or even a single pollinator species are the rule in the family. Shifts in pollination system are more frequent in genera of Crocoideae that have bilaterally symmetric flowers and a perianth tube, features that promote adaptive radiation by facilitating precise shifts in pollen placement, in conjunction with changes in flower colour, scent and tube length.
• Conclusions Diversity of pollination systems explains in part the huge species diversity of Iridaceae in sub-Saharan Africa, and permits species packing locally. Pollination shifts are, however, seen as playing a secondary role in speciation by promoting reproductive isolation in peripheral, ecologically distinct populations in areas of diverse topography, climate and soils. Pollination of Iridaceae in Eurasia and the New World, where the family is also well represented, is poorly studied but appears less diverse, although pollination by both pollen- and oil-collecting bees is frequent and bird pollination rare.
doi:10.1093/aob/mcj040
PMCID: PMC2803647  PMID: 16377653
Floral form; fragrance chemistry; guilds; keystone species; nectar chemistry; Coleoptera; Hymenoptera; Lepidoptera; Nectarinia
4.  Pterandra pyroidea: a case of pollination shift within Neotropical Malpighiaceae 
Annals of Botany  2011;107(8):1323-1334.
Background and Aims
Most Neotropical species of Malpighiaceae produce floral fatty oils in calyx glands to attract pollinating oil-collecting bees, which depend on this resource for reproduction. This specialized type of pollination system tends to be lost in members of the family that occur outside the geographic distribution (e.g. Africa) of Neotropical oil-collecting bees. This study focused on the pollination ecology, chemical ecology and reproductive biology of an oil flower species, Pterandra pyroidea (Malpighiaceae) from the Brazilian Cerrado. Populations of this species consist of plants with oil-secreting (glandular) flowers, plants with non-oil-secreting flowers (eglandular) or a mix of both plant types. This study specifically aims to clarify the role of eglandular morphs in this species.
Methods
Data on pollinators were recorded by in situ observations. Breeding system experiments were conducted by isolating inflorescences and by enzymatic reactions. Floral resources, pollen and floral oils offered by this species were analysed by staining and a combination of various spectroscopic methods.
Key Results
Eglandular flowers of P. pyroidea do not act as mimics of their oil-producing conspecifics to attract pollinators. Instead, both oil-producing and oil-free flowers depend on pollen-collecting bees for reproduction, and their main pollinators are bumble-bees. Floral oils produced by glandular flowers are less complex than those described in closely related genera.
Conclusions
Eglandular flowers represent a shift in the pollination system in which oil is being lost and pollen is becoming the main reward of P. pyroidea flowers. Pollination shifts of this kind have hitherto not been demonstrated empirically within Neotropical Malpighiaceae and this species exhibits an unusual transition from a specialized towards a generalized pollination system in an area considered the hotspot of oil-collecting bee diversity in the Neotropics. Transitions of this type provide an opportunity to study ongoing evolutionary mechanisms that promote the persistence of species previously involved in specialized mutualistic relationships.
doi:10.1093/aob/mcr084
PMCID: PMC3101150  PMID: 21610210
Cerrado; elaiophores; fatty acids; floral oils; floral rewards; gas chromatography; Malpighiaceae; oil-collecting bees; oil flowers; pollination shift; pollination syndromes; Pterandra pyroidea
5.  A pollinators' eye view of a shelter mimicry system 
Annals of Botany  2013;111(6):1155-1165.
Background and Aims
‘Human-red’ flowers are traditionally considered to be rather unpopular with bees, yet some allogamous species in the section Oncocyclus (genus Iris, Iridaceae) have evolved specialized interactions with their pollinators, a narrow taxonomic range of male solitary bees. The dark-red, tubular flowers of these irises are nectarless but provide protective shelters (i.e. a non-nutritive form of reward) primarily to male solitary bees (Apidae, Eucerini) that pollinate the flowers while looking for a shelter. An earlier study on orchids suggested that species pollinated predominantly by male solitary bees produce significantly larger amounts and larger numbers of different n-alkenes (unsaturated cuticular hydrocarbons). Whether or not this also applies to the Oncocyclus irises and whether pollinators are attracted by specific colours or scents of these flowers is unknown.
Methods
Using Iris atropurpurea, recording of pollinator preferences for shelters with different spatial parameters was combined with analyses of floral colours (by spectrophotometry) and scents (by gas chromatography–mass spectrometry) to test the hypotheses that (a) pollinators significantly prefer floral tunnels facing the rising sun (floral heat-reward hypothesis), and that (b) flowers pollinated predominantly by male solitary bees produce significantly larger amounts and larger numbers of unsaturated cuticular hydrocarbons (n-alkenes) in their floral scent (preadaptation to sexual-deception hypothesis).
Key Results
Male bees do not significantly prefer shelters facing the rising sun or with the presence of high absolute/relative amounts and numbers of n-alkenes in the floral scent.
Conclusions
The results suggest that the flowers of I. atropurpurea probably evolved by pollinator-mediated selection acting primarily on floral colours to mimic large achromatic (‘bee-black’) protective shelters used preferentially by male solitary bees, and that pollinator visits are presumably not the result of an odour-based sexual stimulation or motivated by an increased morning floral heat reward in tunnels facing the rising sun.
doi:10.1093/aob/mct081
PMCID: PMC3662522  PMID: 23599249
Shelter mimicry; floral evolution; pollinator preferences; floral scents; floral colours; Iris atropurpurea; Oncocyclus
6.  Floral elaiophore structure in four representatives of the Ornithocephalus clade (Orchidaceae: Oncidiinae) 
Annals of Botany  2012;110(4):809-820.
Background and Aims
A significant number of species assigned to the Neotropical orchid sub-tribe Oncidiinae reward insect pollinators with oil produced in floral glands termed elaiophores. The latter may be glabrous (epithelial elaiophores) or hirsute (trichomal elaiophores). Although the detailed anatomy and ultrastructure of epithelial elaiophores have been studied for a number of genera, such as Oncidium Sw., Gomesa R. Br. and Trichocentrum Poepp. & Endl., hitherto, trichomal elaiophores have been investigated only for a single species of Oncidiinae, Ornithocephalus ciliatus Lindl. Furthermore, this is the only representative of the Ornithocephalus clade to be investigated to date. Here, an examination is made of the elaiophore anatomy and ultrastructure of a further four species currently assigned to this clade (Ornithocephalus gladiatus Hook., Phymatidium falcifolium Lindl., Zygostates grandiflora (Lindl.) Mansf. and Zygostates lunata Lindl.) and the results compared with those obtained for other Oncidiinae.
Methods
Elaiophore structure was examined for all species at three stages of flower development: closed bud, first day of anthesis and final stage of anthesis, using light microscopy, fluorescence microscopy, scanning electron microscopy, transmission electron microscopy and histochemistry.
Key Results
Elaiophores of O. gladiatus occur upon the lateral lobes of the labellum and display characters intermediate between those of typical epithelial and trichomal elaiophores, in that they are largely glabrous, consisting mainly of cuboidal epidermal cells, but bear short, unicellular hairs proximally. By contrast, the elaiophores of all the other species investigated occur on the callus and are of the trichomal type. In P. falcifolium, these unicellular hairs are capitate. In all species, oil secretion commenced at the closed floral bud stage. Ultrastructurally, the mainly trichomal elaiophores of the four representatives of the Ornithocephalus clade closely resembled the epithelial elaiophores of other Oncidiinae, in that their cells displayed an organelle complement typical of lipid-secreting cells. However, in some taxa, a number of noteworthy characters were present. For example, the elaiophore cuticle of O. gladiatus and P. falcifolium was bi-layered, the outer layer being lamellate, the inner reticulate. The cuticle of Z. grandiflora and Z. lunata was also lamellate, but here, a reticulate layer was absent. Accumulation of secreted oil resulted in the localized distension of the cuticle. Cuticular cracks and pores, however, were absent from all species. The walls of the secretory cells of Z. grandiflora were also atypical in that they had short protuberances or ingrowths, and contained cavities which are thought to be involved in the secretory process.
Conclusions
Of the species investigated, most displayed similar anatomical organization, their trichomal elaiophores occurring on the labellar callus. They, thus, differ from many other members of the Oncidiinae, where epithelial elaiophores are found either on the callus, or on the lateral lobes of the labellum. However, ultrastructurally, all elaiophores, whether those of representatives of the Ornithocephalus clade, or those of other oil-secreting Oncidiinae, possessed a similar complement of organelles, regardless of whether the elaiophores were trichomal or epithelial. In view of the latter, and the similar chemical composition of oils derived from all Oncidiinae investigated to date, it is probable that position and type of elaiophore, and possibly the structure of the overlying cuticle, play an important role in pollinator selection in these oil-secreting orchids.
doi:10.1093/aob/mcs158
PMCID: PMC3423815  PMID: 22805528
Anatomy; elaiophore; histochemistry; lipid secretion; micromorphology; oil glands; Oncidiinae; trichome; ultrastructure
7.  Floral and vegetative cues in oil-secreting and non-oil-secreting Lysimachia species 
Annals of Botany  2012;110(1):125-138.
Background and Aims
Unrelated plants pollinated by the same group or guild of animals typically evolve similar floral cues due to pollinator-mediated selection. Related plant species, however, may possess similar cues either as a result of pollinator-mediated selection or as a result of sharing a common ancestor that possessed the same cues or traits. In this study, visual and olfactory floral cues in Lysimachia species exhibiting different pollination strategies were analysed and compared, and the importance of pollinators and phylogeny on the evolution of these floral cues was determined. For comparison, cues of vegetative material were examined where pollinator selection would not be expected.
Methods
Floral and vegetative scents and colours in floral oil- and non-floral oil-secreting Lysimachia species were studied by chemical and spectrophotometric analyses, respectively, compared between oil- and non-oil-secreting species, and analysed by phylogenetically controlled methods.
Key Results
Vegetative and floral scent was species specific, and variability in floral but not vegetative scent was lower in oil compared with non-oil species. Overall, oil species did not differ in their floral or vegetative scent from non-oil species. However, a correlation was found between oil secretion and six floral scent constituents specific to oil species, whereas the presence of four other floral compounds can be explained by phylogeny. Four of the five analysed oil species had bee-green flowers and the pattern of occurrence of this colour correlated with oil secretion. Non-oil species had different floral colours. The colour of leaves was similar among all species studied.
Conclusions
Evidence was found for correlated evolution between secretion of floral oils and floral but not vegetative visual and olfactory cues. The cues correlating with oil secretion were probably selected by Macropis bees, the specialized pollinators of oil-secreting Lysimachia species, and may have evolved in order to attract these bees.
doi:10.1093/aob/mcs101
PMCID: PMC3380597  PMID: 22634256
Colour hexagon; oil secretion; correlated evolution; flower and vegetative scent; headspace analysis; GC-MS; Lysimachia; multidimensional scaling; oil-bee Macropis; phylogeny; spectral photometry
8.  The evolution and loss of oil-offering flowers: new insights from dated phylogenies for angiosperms and bees 
The interactions between bees that depend on floral oil for their larvae and flowers that offer oil involve an intricate mix of obligate and facultative mutualisms. Using recent phylogenies, new data on oil-offering Cucurbitaceae, and molecular-dating, we ask when and how often oil-offering flowers and oil-foraging bees evolved, and how frequently these traits were lost in the cause of evolution. Local phylogenies and an angiosperm-wide tree show that oil flowers evolved at least 28 times and that floral oil was lost at least 36–40 times. The oldest oil flower systems evolved shortly after the K/T boundary independently in American Malpighiaceae, tropical African Cucurbitaceae and Laurasian Lysimachia (Myrsinaceae); the ages of the South African oil flower/oil bee systems are less clear. Youngest oil flower clades include Calceolaria (Calceolariaceae), Iridaceae, Krameria (Krameriaceae) and numerous Orchidaceae, many just a few million years old. In bees, oil foraging evolved minimally seven times and dates back to at least 56 Ma (Ctenoplectra) and 53 Ma (Macropis). The co-occurrence of older and younger oil-offering clades in three of the four geographical regions (but not the Holarctic) implies that oil-foraging bees acquired additional oil hosts over evolutionary time. Such niche-broadening probably started with exploratory visits to flowers resembling oil hosts in scent or colour, as suggested by several cases of Muellerian or Batesian mimicry involving oil flowers.
doi:10.1098/rstb.2009.0229
PMCID: PMC2838259  PMID: 20047869
oil-offering flowers; oil-foraging bees; molecular clock dating; evolutionary gain; evolutionary loss; oil biochemistry
9.  Fly pollination in Ceropegia (Apocynaceae: Asclepiadoideae): biogeographic and phylogenetic perspectives 
Annals of Botany  2009;103(9):1501-1514.
Background and Aims
Ceropegia (Apocynaceae subfamily Asclepiadoideae) is a large, Old World genus of >180 species, all of which possess distinctive flask-shaped flowers that temporarily trap pollinators. The taxonomic diversity of pollinators, biogeographic and phylogenetic patterns of pollinator exploitation, and the level of specificity of interactions were assessed in order to begin to understand the role of pollinators in promoting diversification within the genus.
Methods
Flower visitor and pollinator data for approx. 60 Ceropegia taxa were analysed with reference to the main centres of diversity of the genus and to a cpDNA–nrDNA molecular phylogeny of the genus.
Key Results
Ceropegia spp. interact with flower-visiting Diptera from at least 26 genera in 20 families, of which 11 genera and 11 families are pollinators. Size range of flies was 0·5–4·0 mm and approx. 94 % were females. Ceropegia from particular regions do not use specific fly genera or families, though Arabian Peninsula species are pollinated by a wider range of Diptera families than those in other regions. The basal-most clade interacts with the highest diversity of Diptera families and genera, largely due to one hyper-generalist taxon, C. aristolochioides subsp. deflersiana. Species in the more-derived clades interact with a smaller diversity of Diptera. Approximately 60 % of taxa are so far recorded as interacting with only a single genus of pollinators, the remaining 40 % being less conservative in their interactions. Ceropegia spp. can therefore be ecological specialists or generalists.
Conclusions
The genus Ceropegia has largely radiated without evolutionary shifts in pollinator functional specialization, maintaining its interactions with small Diptera. Intriguing biogeographic and phylogenetic patterns may reflect processes of regional dispersal, diversification and subsequent specialization onto a narrower range of pollinators, though some of the findings may be caused by inconsistent sampling. Comparisons are made with other plant genera in the Aristolochiaceae and Araceae that have evolved flask-shaped flowers that trap female flies seeking oviposition sites.
doi:10.1093/aob/mcp072
PMCID: PMC2701756  PMID: 19339298
Apocynaceae; Asclepiadoideae; Brachystelma; Ceropegia; Diptera; flower evolution; generalization; mutualism; pollination; Riocreuxia; specialization; Stapeliinae
10.  Studies on diversity and evolution of Iridaceae species in southern Brazil 
Genetics and Molecular Biology  2012;35(4 Suppl):1027-1035.
Plants of the family Iridaceae are well represented in the grassland vegetation of southern Brazil, occurring in the Pampa and Atlantic Forest biomes. Nevertheless, little is known about the taxonomy and evolution of Iridaceae species in southern Brazil. The main goal of this review is to compile published information about South American Iridaceae, and to discuss the evolution and genetic diversity of the family presenting our own research data in the light of the published literature. The main focus is on the genera Calydorea, Cypella, Herbertia, and Sisyrinchium. Aspects of reproductive system and of pollinator attraction are also discussed.
PMCID: PMC3571435  PMID: 23412701
cytotaxonomy; molecular phylogenetics; Iridoideae; population genetics; diversity
11.  The endangered Iris atropurpurea (Iridaceae) in Israel: honey-bees, night-sheltering male bees and female solitary bees as pollinators 
Annals of Botany  2012;111(3):395-407.
Background and Aims
The coastal plain of Israel hosts the last few remaining populations of the endemic Iris atropurpurea (Iridaceae), a Red List species of high conservation priority. The flowers offer no nectar reward. Here the role of night-sheltering male solitary bees, honey-bees and female solitary bees as pollinators of I. atropurpurea is documented.
Methods
Breeding system, floral longevity, stigma receptivity, visitation rates, pollen loads, pollen deposition and removal and fruit- and seed-set were investigated.
Key Results
The main wild pollinators of this plant are male eucerine bees, and to a lesser extent, but with the potential to transfer pollen, female solitary bees. Honey-bees were found to be frequent diurnal visitors; they removed large quantities of pollen and were as effective as male sheltering bees at pollinating this species. The low density of pollen carried by male solitary bees was attributed to grooming activities, pollen displacement when bees aggregated together in flowers and pollen depletion by honey-bees. In the population free of honey-bee hives, male bees carried significantly more pollen grains on their bodies. Results from pollen analysis and pollen deposited on stigmas suggest that inadequate pollination may be an important factor limiting fruit-set. In the presence of honey-bees, eucerine bees were low removal–low deposition pollinators, whereas honey-bees were high removal–low deposition pollinators, because they removed large amounts into corbiculae and deposited relatively little onto receptive stigmas.
Conclusions
Even though overall, both bee taxa were equally effective pollinators, we suggest that honey-bees have the potential to reduce the amount of pollen available for plant reproduction, and to reduce the amount of resources available to solitary bee communities. The results of this study have potential implications for the conservation of this highly endangered plant species if hives are permitted inside reserves, where the bulk of Oncocyclus iris species are protected.
doi:10.1093/aob/mcs292
PMCID: PMC3579445  PMID: 23275630
Endangered; Iris atropurpurea; pollination; pollinator effectiveness; Apis mellifera; night-sheltering; eucerine bees; solitary bees; pollen removal; pollen deposition; stigma receptivity; pollen viability
12.  Floral convergence in Oncidiinae (Cymbidieae; Orchidaceae): an expanded concept of Gomesa and a new genus Nohawilliamsia 
Annals of Botany  2009;104(3):387-402.
Background
Floral morphology, particularly the angle of lip attachment to the column, has historically been the fundamental character used in establishing generic limits in subtribe Oncidiinae (Orchidaceae), but it has also been long recognized that reliance on this character alone has produced a highly artificial set of genera. In essence, lip/column relationships reflect syndromes associated with pollinator preferences; most genera of Oncidiinae as previously defined have consisted of a single floral type. Here, the degree to which this has influenced generic delimitation in Brazilian members of the largest genus of Oncidiinae, Oncidium, which previous molecular (DNA) studies have demonstrated to be polyphyletic, is evaluated.
Methods
Phylogenetic analyses of the following multiple DNA regions were used: the plastid psbA-trnH intergenic spacer, matK exon and two regions of ycf1 exon and nuclear ribosomal DNA, comprised of the two internal transcribed spacers, ITS1 and ITS2, and the 5·8S gene. Results from all regions analysed separately indicated highly similar relationships, so a combined matrix was analysed.
Key Results
Nearly all species groups of Brazilian Oncidium are only distantly related to the type species of the genus, O. altissimum, from the Caribbean. There are two exceptions to this geographical rule: O. baueri is related to the type group and O. orthostates, an isolated species that lacks the defining tabula infrastigmata of Oncidium, is not exclusively related to any previously described genus in the subtribe. Several well-supported subclades can be observed in these results, but they do not correspond well to sections of Oncidium as previously circumscribed or to segregate genera as defined by several recent authors. In spite of their floral differences, these groups of Oncidium, formerly treated as O. sections Barbata, Concoloria pro parte, Crispa, Ranifera, Rhinocerotes, Rostrata (only O. venustum), Synsepala, Verrucituberculata pro parte and Waluewa, form a well-supported clade with Gomesa (including Rodrigueziella and Rodrigueziopsis) embedded in it. Two often recognized segregate genera, Baptistonia and Ornithophora, and the recently described Carriella are also embedded within the Brazilian clade. The level of variation within major subclades of the Gomesa clade is low and similar to that observed within other genera of Oncidiinae.
Conclusions
Convergence on a stereotypical syndrome of floral traits associated with pollination by oil-collecting bees has resulted in these characters not being reliable for producing monophyletic taxa, and the genus Oncidium, defined by these characters, is grossly polyphyletic. Vegetative and a few floral/inflorescence characters link these taxa with a mainly Brazilian distribution, and they were all transferred to Gomesa on this basis rather than separated from Gomesa based on their floral differences, which we hypothesize to be simple shifts in pollination strategies. Other authors have described a large number of new genera for these former members of Oncidium, but most of these are not supported by the results presented here (i.e. they are not monophyletic). A new genus, Nohawilliamsia, is described for O. orthostates because it does not fit in any currently recognized genus and is only distantly related to any other member of Oncidiinae.
doi:10.1093/aob/mcp067
PMCID: PMC2720657  PMID: 19346522
Baptistonia; Brazilian orchids; Carriella; deceit pollination; Gomesa; ITS; matK; oil-collecting bees; Oncidium; Oncidiinae; Orchidaceae
13.  Comparative labellar micromorphology of Zygopetalinae (Orchidaceae) 
Annals of Botany  2011;108(5):945-964.
Background and Aims
Molecular evidence indicates that the Neotropical sub-tribe Zygopetalinae is sister to Maxillariinae. Most members of the latter sub-tribe have deceit pollination strategies, but some species produce rewards such as nectar, pseudopollen, resin and wax, and are pollinated by a range of pollinators that include stingless bees (Meliponini), wasps and hummingbirds. By contrast, relatively little is known about the pollination of Zygopetalinae species. However, some are pollinated by fragrance-gathering, male euglossine bees or employ nectar deceit strategies. The aim of this study is to describe the labellar micromorphology of Zygopetalinae and to compare it with that of Maxillariinae sensu lato (s.l.) as part of an ongoing project to record the range of labellar characters found within the tribe Maxillarieae, and to assess whether these characters represent synapomorphies or homoplasies resulting from similar pollination pressures.
Methods
The labella of 31 species of Zygopetalinae, including Cryptarrhena R. Br. and representatives of the Zygopetalum, Huntleya and Warrea clades, were examined using light microscopy and scanning electron microscopy, and the range of labellar characters was recorded. These characters were subsequently compared with those of Maxillariinae s.l. which formed the subject of our previous investigations.
Key Results and Conclusions
The labellar micromorphology of Zygopetalinae is less diverse than that of Maxillariinae and does not reflect the currently accepted phylogeny of the former sub-tribe based on molecular studies. Instead, the relative uniformity in labellar micromorphology of Zygopetalinae is probably due to homoplasies resulting from similar pollinator pressures. Labellar trichomes are relatively uncommon in Zygopetalinae, but occur in certain members of both the Zygopetalum and Huntleya clades. Trichomes are unbranched, uniseriate and multicellular with rounded apices, or unbranched and unicellular, with tapering, pointed and flexuose apices. Hitherto, unicellular trichomes of this kind have been observed only for euglossophilous orchid taxa, and the adoption of a relatively limited range of pollination strategies by Zygopetalinae may have resulted in reduced investment in micromorphological labellar characters.
doi:10.1093/aob/mcr202
PMCID: PMC3177679  PMID: 21856635
Cryptarrhena; epidermis; homoplasy; Huntleya clade; labellum; Maxillariinae; papillae; trichomes; Warrea clade; Zygopetalum clade
14.  Labellar Micromorphology of Two Euglossine-pollinated Orchid Genera; Scuticaria Lindl. and Dichaea Lindl. 
Annals of Botany  2008;102(5):805-824.
Background and Aims
Until recently, there was no consensus regarding the phylogenetic relationships of the Neotropical orchid genera Scuticaria Lindl. and Dichaea Lindl. However, recent evidence derived from both gross morphological and molecular studies supports the inclusion of Scuticaria and Dichaea in sub-tribes Maxillariinae and Zygopetalinae, respectively. The present paper describes the labellar micromorphology of both genera and seeks to establish whether labellar characters support the assignment of Scuticaria and Dichaea to these sub-tribes.
Methods
The labella of four species of Scuticaria and 14 species of Dichaea were examined using light microscopy and scanning electron microscopy, and their micromorphology was compared with that of representative species of Maxillariinae sensu lato and Zygopetalinae (Huntleya clade).
Key Results and Conclusions
In most specimens of Scuticaria examined, the papillose labella bear uniseriate, multicellular, unbranched trichomes. However, in S. steelii (Lindl.) Lindl., branched hairs may also be present and some trichomes may fragment and form pseudopollen. Multicellular, leaf-like scales were also present in one species of Scuticaria. Similar, unbranched hairs are present in certain species of Maxillaria Ruiz & Pav. (Maxillariinae sensu stricto) and Chaubardia Rchb.f. (Huntleya clade). As yet, moniliform, pseudopollen-forming hairs have not been observed for Zygopetalinae, but their presence in Scuticaria steelii, Maxillaria and Heterotaxis Lindl. supports the placing of Scuticaria in Maxillariinae. As other genera are sampled, the presence of branched hairs, hitherto unknown for Maxillariinae sensu lato, may prove to be a useful character in taxonomy and phylogenetic studies. Euglossophily occurs in Dichaea, as well as Chondrorhyncha Lindl. and Pescatorea Rchb.f. (Huntleya clade), and all three genera tend to lack distinctive labellar features. Instead, lip micromorphology is relatively simple and glabrous or papillose. However, two of the Dichaea species examined bear unicellular, labellar trichomes very similar to those found in Bifrenaria Lindl. (pollinated by both euglossine bees and Bombus spp.), and this feature may have arisen by convergence in response to similar pollination pressures.
doi:10.1093/aob/mcn155
PMCID: PMC2712378  PMID: 18765439
Bifrenaria; Bifrenaria clade; Chaubardia; Chondrorhyncha; Dichaea; Dichaeinae; Heterotaxis; Huntleya clade; Huntleyinae; labellum; Maxillaria; Maxillariinae; papillae; Pescatorea; scales; Scuticaria; trichomes; Zygopetalinae
15.  Global characterization of Artemisia annua glandular trichome transcriptome using 454 pyrosequencing 
BMC Genomics  2009;10:465.
Background
Glandular trichomes produce a wide variety of commercially important secondary metabolites in many plant species. The most prominent anti-malarial drug artemisinin, a sesquiterpene lactone, is produced in glandular trichomes of Artemisia annua. However, only limited genomic information is currently available in this non-model plant species.
Results
We present a global characterization of A. annua glandular trichome transcriptome using 454 pyrosequencing. Sequencing runs using two normalized cDNA collections from glandular trichomes yielded 406,044 expressed sequence tags (average length = 210 nucleotides), which assembled into 42,678 contigs and 147,699 singletons. Performing a second sequencing run only increased the number of genes identified by ~30%, indicating that massively parallel pyrosequencing provides deep coverage of the A. annua trichome transcriptome. By BLAST search against the NCBI non-redundant protein database, putative functions were assigned to over 28,573 unigenes, including previously undescribed enzymes likely involved in sesquiterpene biosynthesis. Comparison with ESTs derived from trichome collections of other plant species revealed expressed genes in common functional categories across different plant species. RT-PCR analysis confirmed the expression of selected unigenes and novel transcripts in A. annua glandular trichomes.
Conclusion
The presence of contigs corresponding to enzymes for terpenoids and flavonoids biosynthesis suggests important metabolic activity in A. annua glandular trichomes. Our comprehensive survey of genes expressed in glandular trichome will facilitate new gene discovery and shed light on the regulatory mechanism of artemisinin metabolism and trichome function in A. annua.
doi:10.1186/1471-2164-10-465
PMCID: PMC2763888  PMID: 19818120
16.  An extinct Eocene taxon of the daisy family (Asteraceae): evolutionary, ecological and biogeographical implications 
Annals of Botany  2012;109(1):127-134.
Background and Aims
Morphological, molecular and biogeographical information bearing on early evolution of the sunflower alliance of families suggests that the clade containing the extant daisy family (Asteraceae) differentiated in South America during the Eocene, although palaeontological studies on this continent failed to reveal conclusive support for this hypothesis. Here we describe in detail Raiguenrayun cura gen. & sp. nov., an exceptionally well preserved capitulescence of Asteraceae recovered from Eocene deposits of northwestern Patagonia, Argentina.
Methods
The fossil was collected from the 47·5 million-year-old Huitrera Formation at the Estancia Don Hipólito locality, Río Negro Province, Argentina.
Key Results
The arrangement of the capitula in a cymose capitulescence, the many-flowered capitula with multiseriate–imbricate involucral bracts and the pappus-like structures indicate a close morphological relationship with Asteraceae. Raiguenrayun cura and the associated pollen Mutisiapollis telleriae do not match exactly any living member of the family, and clearly represent extinct taxa. They share a mosaic of morphological features today recognized in taxa phylogenetically close to the root of Asteraceae, such as Stifftieae, Wunderlichioideae and Gochnatieae (Mutisioideae sensu lato) and Dicomeae and Oldenburgieae (Carduoideae), today endemic to or mainly distributed in South America and Africa, respectively.
Conclusions
This is the first fossil genus of Asteraceae based on an outstandingly preserved capitulescence that might represent the ancestor of Mutisioideae–Carduoideae. It might have evolved in southern South America some time during the early Palaeogene and subsequently entered Africa, before the biogeographical isolation of these continents became much more pronounced. The new fossil represents the first reliable point for calibration, favouring an earlier date to the split between Barnadesioideae and the rest of Asteraceae than previously thought, which can be traced back at least 47·5 million years. This is the oldest well dated member of Asteraceae and perhaps the earliest indirect evidence for bird pollination in the family.
doi:10.1093/aob/mcr240
PMCID: PMC3241571  PMID: 22179952
Compositae; capitulescence; fossil taxon; Raiguenrayun cura gen. & sp. nov.; Eocene; Patagonia; southern South America
17.  Response of Sunflower (Helianthus annuus L.) Leaf Surface Defenses to Exogenous Methyl Jasmonate 
PLoS ONE  2012;7(5):e37191.
Helianthus annuus, the common sunflower, produces a complex array of secondary compounds that are secreted into glandular trichomes, specialized structures found on leaf surfaces and anther appendages of flowers. The primary components of these trichome secretions are sesquiterpene lactones (STL), a diverse class of compounds produced abundantly by the plant family Compositae and believed to contribute to plant defense against herbivory. We treated wild and cultivated H. annuus accessions with exogenous methyl jasmonate, a plant hormone that mediates plant defense against insect herbivores and certain classes of fungal pathogens. The wild sunflower produced a higher density of glandular trichomes on its leaves than the cultivar. Comparison of the profiles of glandular trichome extracts obtained by liquid chromatography–mass spectroscopy (LC-MS) showed that wild and cultivated H. annuus were qualitatively similar in surface chemistry, although differing in the relative size and proportion of various compounds detected. Despite observing consistent transcriptional responses to methyl jasmonate treatment, we detected no significant effect on glandular trichome density or LC-MS profile in cultivated or wild sunflower, with wild sunflower exhibiting a declining trend in overall STL production and foliar glandular trichome density of jasmonate-treated plants. These results suggest that glandular trichomes and associated compounds may act as constitutive defenses or require greater levels of stimulus for induction than the observed transcriptional responses to exogenous jasmonate. Reduced defense investment in domesticated lines is consistent with predicted tradeoffs caused by selection for increased yield; future research will focus on the development of genetic resources to explicitly test the ecological roles of glandular trichomes and associated effects on plant growth and fitness.
doi:10.1371/journal.pone.0037191
PMCID: PMC3356381  PMID: 22623991
18.  Evolution of sexual mimicry in the orchid subtribe orchidinae: the role of preadaptations in the attraction of male bees as pollinators 
Background
Within the astonishing diversity of orchid pollination systems, sexual deception is one of the most stunning. An example is the genus Ophrys, where plants attract male bees as pollinators by mimicking female mating signals. Unsaturated hydrocarbons (alkenes) are often the key signal for this chemical mimicry. Here we investigate the evolution of these key compounds within Orchidinae by mapping their production in flowers of selected species onto their estimated phylogeny.
Results
We found that alkenes, at least in trace amounts, were present in 18 of 20 investigated species together representing 10 genera. Thus, the reconstruction of ancestral state for alkene-production showed that this is a primitive character state in Ophrys, and can be interpreted as a preadaptation for the evolution of sexual deception. Four of the investigated species, namely Ophrys sphegodes, Serapias lingua, S. cordigera, and Anacamptis papilionacea, that are pollinated primarily by male bees, produced significantly larger amounts and a greater number of different alkenes than the species pollinated either primarily by female bees or other insects.
Conclusion
We suggest that high amounts of alkenes evolved for the attraction of primarily male bees as pollinators by sensory exploitation, and discuss possible driving forces for the evolution of pollination by male bees.
doi:10.1186/1471-2148-8-27
PMCID: PMC2267782  PMID: 18226206
19.  Glandular Trichomes and Essential Oil of Thymus quinquecostatus 
The Scientific World Journal  2013;2013:387952.
The distribution and types of glandular trichomes and essential oil chemistry of Thymus quinquecostatus were studied. The glandular trichomes are distributed on the surface of stem, leaf, rachis, calyx and corolla, except petiole, pistil and stamen. Three morphologically distinct types of glandular trichomes are described. Peltate trichomes, consisting of a basal cell, a stalk cell and a 12-celled head, are distributed on the stem, leaf, corolla and outer side of calyx. Capitate trichomes, consisting of a unicellular base, a 1–2-celled stalk and a unicellular head, are distributed more diffusely than peltate ones, existing on stem, leaf, rachis and calyx. Digitiform trichomes are just distributed on the outer side of corolla, consisting of 1 basal cell, 3 stalk cells and 1 head cell. All three types of glandular trichomes can secrete essential oil, and in small capitate trichomes of rachis, all peltate trichomes and digitiform trichomes, essential oil is stored in a large subcuticular space, released by cuticle rupture, whereas, in other capitate trichomes, essential oil crosses the thin cuticle. The essential oil of T. quinquecostatus is yellow, and its content is highest in the growth period. 68 constituents were identified in the essential oils. The main constituent is linalool.
doi:10.1155/2013/387952
PMCID: PMC3821910  PMID: 24250266
20.  A comparative survey of floral characters in Capanemia Barb. Rodr. (Orchidaceae: Oncidiinae) 
Annals of Botany  2011;109(1):135-144.
Background and Aims Capanemia
Barb. Rodr. comprises seven species that mostly inhabit the Brazilian Atlantic Rain Forest domain. The genus currently consists of two sections: Capanemia Cogn. and Planifolia Pabst, distinguished on the basis of leaf shape. We compare the floral morphology and anatomy of all species to determine whether separation into sections is supported by floral characters.
Methods
Both fresh flowers and herbarium specimens were investigated, and column and pollinarium features, together with the presence or absence of floral rewards, recorded. Anatomical features were examined using both light microscopy and scanning electron microscopy.
Key Results and Conclusions
With the sole exception of Capanemia therezae, all species shared a distinctive set of floral characters. Flowers were mostly white or yellowish-white and fragrant, and column wings were positioned parallel to the labellum, concealing the stigmatic cavity. Pollinaria had proportionally long tegular stipes and clavate to reniform pollinia, whereas the labellum possessed a conspicuous indument of trichomes, but was devoid of nectar or any other secretion that might function as a food-reward. Capanemia therezae, however, was exceptional in having greenish, unscented flowers with short, rounded and divergent column wings and an exposed stigmatic cavity. Its pollinaria had proportionally short tegular stipes and round pollinia, whereas the labellum lacked trichomes. Droplets of nectar were evident on the adaxial surface of the labellum, adjacent to the callus. Floral features did not support the currently accepted sectional division of Capanemia. If ongoing phylogenetic studies demonstrate that both sections are indeed monophyletic, then these taxa should be distinguished solely on the basis of foliar features.
doi:10.1093/aob/mcr241
PMCID: PMC3241572  PMID: 21937482
Anatomy; column; labellum; morphology; nectary; orchids; pollinarium; pollination; trichomes
21.  The evolution of floral deception in Epipactis veratrifolia (Orchidaceae): from indirect defense to pollination 
BMC Plant Biology  2014;14:63.
Background
It is estimated that floral deception has evolved in at least 7500 species of angiosperms, of which two thirds are orchids. Epipactis veratrifolia (Orchidaceae) is a model system of aphid mimicry as aphidophagous hoverflies lay eggs on false brood sites on their flowers. To understand the evolutionary ecology of floral deception, we investigated the pollination biology of E. veratrifolia across 10 populations in the Eastern Himalayas. We reconstructed the phylogeny of Epipactis and mapped the known pollination systems of previously studied species onto the tree.
Results
Some inflorescences of E. veratrifolia were so infested with aphids while they were still in bud that the some larvae of hoverflies developed to the third instar while flower buds opened. This indicated that adult female hoverflies were partly rewarded for oviposition. Although flowers failed to secrete nectar, they mimicked both alarm pheromones and aphid coloring of to attract female hoverflies as their exclusive pollinators. Phylogenetic mapping indicate that pollination by aphidophagous hoverflies is likely an ancestral condition in the genus Epipactis. We suggest that the biological interaction of aphid (prey), orchid (primary producer) and hoverfly (predator) may represent an intermediate stage between mutualism and deception in the evolution of pollination-by-deceit in E. veratrifolia.
Conclusions
Our analyses indicate that this intermediate stage may be used as a model system to interpret the origin of oviposition (brood site) mimicry in Epipactis. We propose the hypothesis that some deceptive pollination systems evolved directly from earlier (partly) mutualistic systems that maintained the fidelity of the original pollinator(s) even though rewards (nectar/ brood site) were lost.
doi:10.1186/1471-2229-14-63
PMCID: PMC4007573  PMID: 24621377
Anther cap; Aphids; Floral mimicry; Hoverflies; Intermediate stage; Pollinator
22.  Secondary Evolution of a Self-Incompatibility Locus in the Brassicaceae Genus Leavenworthia 
PLoS Biology  2013;11(5):e1001560.
Self-incompatibility enables plants to avoid inbreeding by self-pollination. Here we report that the genetic locus encoding self-pollen recognition has evolved twice in the Brassicaceae family, challenging the notion that loss of self-incompatibility is irreversible.
Self-incompatibility (SI) is the flowering plant reproductive system in which self pollen tube growth is inhibited, thereby preventing self-fertilization. SI has evolved independently in several different flowering plant lineages. In all Brassicaceae species in which the molecular basis of SI has been investigated in detail, the product of the S-locus receptor kinase (SRK) gene functions as receptor in the initial step of the self pollen-rejection pathway, while that of the S-locus cysteine-rich (SCR) gene functions as ligand. Here we examine the hypothesis that the S locus in the Brassicaceae genus Leavenworthia is paralogous with the S locus previously characterized in other members of the family. We also test the hypothesis that self-compatibility in this group is based on disruption of the pollen ligand-producing gene. Sequence analysis of the S-locus genes in Leavenworthia, phylogeny of S alleles, gene expression patterns, and comparative genomics analyses provide support for both hypotheses. Of special interest are two genes located in a non-S locus genomic region of Arabidopsis lyrata that exhibit domain structures, sequences, and phylogenetic histories similar to those of the S-locus genes in Leavenworthia, and that also share synteny with these genes. These A. lyrata genes resemble those comprising the A. lyrata S locus, but they do not function in self-recognition. Moreover, they appear to belong to a lineage that diverged from the ancestral Brassicaceae S-locus genes before allelic diversification at the S locus. We hypothesize that there has been neo-functionalization of these S-locus-like genes in the Leavenworthia lineage, resulting in evolution of a separate ligand-receptor system of SI. Our results also provide support for theoretical models that predict that the least constrained pathway to the evolution of self-compatibility is one involving loss of pollen gene function.
Author Summary
Self-incompatibility (SI) is a pollen recognition system that enables plants to avoid the inbreeding caused by self-pollination. It involves a pair of tightly linked genes known as the S locus. The product of one of these genes acts as the receptor and recognizes the pollen protein produced by the same plant, while the product of the other gene is the pollen protein that is recognized by the receptor. In this study, we have analyzed the gene sequence, genome organization, and gene evolutionary history of S loci in members of the Brassicaceae family, which includes plants of the genus Leavenworthia. From our analyses, we conclude that both genes that comprise the ancestral S locus in the Brassicaceae were lost in Leavenworthia. We show, however, that plants of this genus possess two other linked genes that exhibit patterns of polymorphism and expression that are characteristic of an S locus. These genes occupy the same genomic position in Leavenworthia as do two non-S-locus genes in the related species Arabidopsis lyrata, genes that are not known to function in self-recognition in this species. We suggest that these genes have evolved to assume the function of the pollen recognition system of SI in Leavenworthia—that is, that there has been de novo emergence of a distinct Brassicaceae S locus in this genus. We also present evidence that the breakdown of the SI system in two Leavenworthia races is due to independent mutations in the S-locus pollen gene, in accordance with theoretical predictions for the spread of S-locus disrupting mutations.
doi:10.1371/journal.pbio.1001560
PMCID: PMC3653793  PMID: 23690750
23.  Floral features, pollination biology and breeding system of Chloraea membranacea Lindl. (Orchidaceae: Chloraeinae) 
Annals of Botany  2012;110(8):1607-1621.
Background and Aims
The pollination biology of very few Chloraeinae orchids has been studied to date, and most of these studies have focused on breeding systems and fruiting success. Chloraea membranacea Lindl. is one of the few non-Andean species in this group, and the aim of the present contribution is to elucidate the pollination biology, functional floral morphology and breeding system in native populations of this species from Argentina (Buenos Aires) and Brazil (Rio Grande do Sul State).
Methods
Floral features were examined using light microscopy, and scanning and transmission electron microscopy. The breeding system was studied by means of controlled pollinations applied to plants, either bagged in the field or cultivated in a glasshouse. Pollination observations were made on natural populations, and pollinator behaviour was recorded by means of photography and video.
Key Results
Both Argentinean and Brazilian plants were very consistent regarding all studied features. Flowers are nectarless but scented and anatomical analysis indicates that the dark, clavate projections on the adaxial labellar surface are osmophores (scent-producing glands). The plants are self-compatible but pollinator-dependent. The fruit-set obtained through cross-pollination and manual self-pollination was almost identical. The main pollinators are male and female Halictidae bees that withdraw the pollinarium when leaving the flower. Remarkably, the bees tend to visit more than one flower per inflorescence, thus promoting self-pollination (geitonogamy). Fruiting success in Brazilian plants reached 60·78 % in 2010 and 46 % in 2011. Some pollinarium-laden female bees were observed transferring pollen from the carried pollinarium to their hind legs. The use of pollen by pollinators is a rare record for Orchidaceae in general.
Conclusions
Chloraea membrancea is pollinated by deceit. Together, self-compatibility, pollinarium texture, pollinator abundance and behaviour may account for the observed high fruiting success. It is suggested that a reappraisal and re-analysis of important flower features in Chloraeinae orchids is necessary.
doi:10.1093/aob/mcs221
PMCID: PMC3503500  PMID: 23071217
Breeding system; Chloraea membrancea; Chloraeinae; Halictidae; Orchidaceae; orchids; pollination
24.  Effective pollinators of Asian sacred lotus (Nelumbo nucifera): contemporary pollinators may not reflect the historical pollination syndrome 
Annals of Botany  2009;104(5):845-851.
Background and Aims
If stabilizing selection by pollinators is a prerequisite for pollinator-mediated floral evolution, spatiotemporal variation in the pollinator assemblage may confuse the plant–pollinator interaction in a given species. Here, effective pollinators in a living fossil plant Nelumbo nucifera (Nelumbonaceae) were examined to test whether beetles are major pollinators as predicted by its pollination syndrome.
Methods
Pollinators of N. nucifera were investigated in 11 wild populations and one cultivated population, and pollination experiments were conducted to examine the pollinating role of two major pollinators (bees and beetles) in three populations.
Key Results
Lotus flowers are protogynous, bowl shaped and without nectar. The fragrant flowers can be self-heating during anthesis and produce around 1 million pollen grains per flower. It was found that bees and flies were the most frequent flower visitors in wild populations, contributing on average 87·9 and 49·4 % of seed set in Mishan and Lantian, respectively. Beetles were only found in one wild population and in the cultivated population, but the pollinator exclusion experiments showed that beetles were effective pollinators of Asian sacred lotus.
Conclusions
This study indicated that in their pollinating role, beetles, probable pollinators for this thermoregulating plant, had been replaced by some generalist insects in the wild. This finding implies that contemporary pollinators may not reflect the pollination syndrome.
doi:10.1093/aob/mcp173
PMCID: PMC2749538  PMID: 19617594
Nelumbo nucifera; beetle pollinated; pollination syndrome; effective pollinator; generalized flower; thermoregulation; Nelumbonaceae
25.  Labellar Micromorphology of Bifrenariinae Dressler (Orchidaceae) 
Annals of Botany  2006;98(6):1215-1231.
• Background and Aims The two closely related subtribes Bifrenariinae Dressler and Maxillariinae Benth. are easily distinguished on morphological grounds. Recently, however, molecular techniques have supported the inclusion of Bifrenariinae within a more broadly defined Maxillariinae. The present paper describes the diverse labellar micromorphology found amongst representatives of Bifrenariinae (Bifrenaria Lindl., Rudolfiella Hoehne, Teuscheria Garay and Xylobium Lindl.) and compares it with that found in Maxillaria Pabst & Dungs and Mormolyca Fenzl (Maxillariinae).
• Methods The labella of 35 specimens representing 22 species of Bifrenariinae were examined by means of light microscopy and scanning electron microscopy and their micromorphology compared with that of Maxillaria sensu stricto and Mormolyca spp. The labellar epidermis of representatives of Bifrenaria, Xylobium and Mormolyca was tested for protein, starch and lipids in order to ascertain whether this tissue is involved in the rewarding of pollinators.
• Key Results and Conclusions The labella of Bifrenaria spp. and Mormolyca spp. are densely pubescent but those of Xylobium, Teuscheria and Rudolfiella are generally papillose. However, whereas the trichomes of Bifrenaria and Mormolyca are unicellular, those found in the other three genera are multicellular. Hitherto, no unicellular trichomes have been described for Maxillaria, although the labella of a number of species secrete a viscid substance or bear moniliform, pseudopollen-producing hairs. Moniliform hairs and secretory material also occur in certain species of Xylobium and Teuscheria and these genera, together with Maxillaria, are thought to be pollinated by stingless bees (Meliponini). Differences in the labellar micromorphology of Bifrenaria and Mormolyca are perhaps related to Euglossine- and/ or bumble bee-mediated pollination and pseudocopulation, respectively. Although Xylobium and Teuscheria share a number of labellar features with Maxillaria sensu stricto, this does not necessarily reflect taxonomic relationships but may be indicative of convergence in response to similar pollinator pressures.
doi:10.1093/aob/mcl204
PMCID: PMC2803581  PMID: 17008352
Bifrenaria; Bifrenariinae; Maxillaria; Maxillariinae; Meliponini; papillae; pollination; pseudopollen; Rudolfiella; Teuscheria; trichomes; Xylobium

Results 1-25 (366514)